Circuit theory emphasizes the importance of edge-crossing decisions in dispersal-scale movements of a forest passerine

Measuring landscape connectivity in ways that reflect an animal’s propensity or reluctance to move across a given landscape is key for planning effective conservation strategies. Resistance distance, based on circuit theory, is one such measure relevant for modeling how broad-scale animal movements over long time periods may lead to gene flow across the landscape. Despite the success of circuit theory in landscape genetic studies, its applicability to model finer-scale processes such as the movement patterns of individual animals within their breeding grounds (e.g., while prospecting for territories) has yet to be tested. Here, we applied both circuit models and least-cost models to understand the relationship between landscape connectivity and return time of Ovenbirds (Seiurus aurocapilla) that had been translocated at least 20 km from their home territory near Québec City, Canada. Using an iterative optimization process, we derived resistance values for three cover types (forest, edge, and open) that resulted in resistance distance values that best explained Ovenbird return times. We also identified the cover-type resistance values that yielded length of least-cost path estimates that best explained return times of the translocated birds. The circuit theory and least-cost path methods were equally supported by the data despite being based on different sets of resistance values. The optimal resistance values for calculating resistance distance indicated that for Ovenbirds, traversing a given distance of edge habitat presented a substantially greater resistance than that of open areas. On the other hand, optimized resistances of edge and open were very similar for calculating length of least-cost path. The circuit theory approach suggested that for an Ovenbird moving through fragmented habitat, the number of forest-open transitions (i.e., edge-crossings) that an individual must make is critical to understanding return times after translocation. The least-cost path approach, on the other hand, suggested that the birds strongly avoid all open areas, regardless of size. Circuit theory offers an important new approach for understanding landscapes from the perspective of individuals moving within their breeding range, at finer spatial scales and shorter time scales than have been previously considered.     

Modeling the invasive emerald ash borer risk of spread using a spatially explicit cellular model

The emerald ash borer (EAB, Agrilus planipennis) is decimating native ashes (Fraxinus sp.) throughout midwestern North America, killing millions of trees over the years. With plenty of ash available throughout the continent, the spread of this destructive insect is likely to continue. We estimate that the insect has been moving along a “front” at about 20 km/year since about 1998, but more alarming is its long-range dispersal into new locations facilitated by human activities. We describe a spatially explicit cell-based model used to calculate risk of spread in Ohio, by combining the insect’s flight and short-range dispersal (“insect flight”) with human-facilitated, long-range dispersal (“insect ride”). This hybrid model requires estimates of EAB abundance, ash abundance, major roads and traffic density, campground size and usage, distance from the core infested zone, wood products industry size and type of wood usage, and human population density. With the “insect flight” model, probability of movement is dependent on EAB abundance in the source cells, the quantity of ash in the target cells, and the distances between them. With the “insect-ride” model, we modify the value related to ash abundance based on factors related to potential human-assisted movements of EAB-infested ash wood or just hitchhiking insects. We attempt to show the advantage of our model compared to statistical approaches and to justify its practical value to field managers working with imperfect knowledge. We stress the importance of the road network in distributing insects to new geographically dispersed sites in Ohio, where 84% were within 1 km of a major highway.     

Edge effect and matrix influence on the nest survival of an old forest specialist,the Brown Creeper (<Emphasis Type="Italic">Certhia americana</Emphasis>)

Conservation strategies should be based on a solid understanding of processes underlying species response to landscape change. In forests fragmented by agriculture, elevated nest predation rates have been reported in many forest bird species, especially near edges. In intensively-managed forest landscapes, timber harvesting might also be associated with negative edge effects or broader “context” effects on some species when the matrix provides additional resources to their major nest predators. In this study, we hypothesized that proximity to a forest edge and proportion of cone-producing plantations will increase nest predation risk in fragments of relatively undisturbed forest. We focused on the Brown Creeper (Certhia americana), an indicator species of late-seral forests. We compared habitat configuration and composition at four spatial scales (0.14, 0.5, 1 and 2 km) around 54 nests and related daily nest survival rate to the distance to the nearest forest edge, mean patch size of late-seral forest (r = 141 m), proportion of non-forested lands (r = 141 m), density of maintained roads (r = 1 km), proportion of cone-producing spruce plantations (r = 2 km), and year. The best model included distance to the nearest edge and proportion of cone-producing plantations. Distance of nests to the nearest edge was the best individual predictor of daily nest survival. A larger sample of nests showed a significant threshold in distance to the nearest forest edge; nests located at least 100 m away were more likely to fledge young. These results suggest that even in managed forest landscapes, matrix effects can be important and some bird species may exhibit negative edge effects.     

Landscape context and selection for forest edge by breeding Brown-headed Cowbirds

We evaluated support for four alternate hypotheses explaining the distribution of breeding Brown-headed Cowbirds (Molothrus ater) in forests at varying distances from the forest edge in three Midwestern USA landscapes with varying amounts of forest fragmentation (core forest area ranged from 5 to 70%). We focused on breeding cowbirds’ use of forest because of the risk of nest parasitism to forest-dwelling hosts and to identify factors affecting breeding cowbird habitat selection. We compared distances of cowbird locations in the forest from the forest edge (“edge distances”) to distances of random forest locations in the entire landscape or within individual cowbird home ranges. We analyzed 1322 locations of 84 cowbirds across three landscapes. We found support for the landscape context hypothesis that breeding cowbird preference for forest edge varied with landscape context. Ninety percent of cowbird locations were within 150–350 m of forest edge, despite the overall availability of forest at greater distances from edge (as far as 500–1450 m) both within cowbird home ranges and the entire forested landscape. Cowbird preference for edge varied by landscape context largely due to differences in the availability of forest edge. In a highly fragmented forest cowbirds utilized the entire forest and likely viewed it as “all edge.” In less fragmented forests, cowbirds preferred edge. We consider how variation in cowbird edge preference might relate to patterns in host abundance, host diversity, and host quality because cowbird movements indicate they are capable of using forest farther from edges.     

Maintaining or restoring connectivity of modified landscapes: evaluating the least-cost path model with multiple sources of ecological information

Habitat connectivity is an important element of functioning landscapes for mobile organisms. Maintenance or creation of movement corridors is one conservation strategy for reducing the negative effects of habitat fragmentation. Numerous spatial models exist to predict the location of movement corridors. Few studies, however, have investigated the effectiveness of these methods for predicting actual movement paths. We used an expert-based model and a resource selection function (RSF) to predict least-cost paths of woodland caribou. Using independent data for model evaluation, we found that the expert-based model was a poor predictor of long-distance animal movements; in comparison, the RSF model was effective at predicting habitat selection by caribou. We used the Path Deviation Index (PDI), cumulative path cost, and sinuosity to quantitatively compare the spatial differences between inferred caribou movement paths and predicted least-cost paths, and quasi-random null models of directional movement. Predicted movement paths were on average straighter than inferred movement paths for collared caribou. The PDI indicated that the least-cost paths were no better at predicting the inferred paths than either of two null models—straight line paths and randomly generated paths. We found statistically significant differences in cumulative cost scores for the main effects of model and path type; however, post-hoc comparisons were non-significant suggesting no difference among inferred, random, and predicted least cost paths. Paths generated from an expert based cost surface were more sinuous than those premised on the RSF model, but neither differed from the inferred path. Although our results are specific to one species, they highlight the importance of model evaluation when planning for habitat connectivity. We recommend that conservation planners adopt similar techniques when validating the effectiveness of movement corridors for other populations and species.     

Beyond the least-cost path: evaluating corridor redundancy using a graph-theoretic approach

The impact of the landscape matrix on patterns of animal movement and population dynamics has been widely recognized by ecologists. However, few tools are available to model the matrix’s influence on the length, relative quality, and redundancy of dispersal routes connecting habitat patches. Many GIS software packages can use land use/land cover maps to identify the route of least resistance between two points—the least-cost path. The limitation of this type of analysis is that only a single path is identified, even though alternative paths with comparable costs might exist. In this paper, we implemented two graph theory methods that extend the least-cost path approach: the Conditional Minimum Transit Cost (CMTC) tool and the Multiple Shortest Paths (MSPs) tool. Both methods enable the visualization of multiple dispersal routes that, together, are assumed to form a corridor. We show that corridors containing alternative dispersal routes emerge when favorable habitat is randomly distributed in space. As clusters of favorable habitat start forming, corridors become less redundant and dispersal bottlenecks become visible. Our approach is illustrated using data from a real landscape in the Brazilian Atlantic forest. We explored the effect of small, localized disturbance on dispersal routes linking conservation units. Simulated habitat destruction caused the appearance of alternative dispersal routes, or caused existing corridors to become narrower. These changes were observed even in the absence of significant differences in the length or cost of least-cost paths. Last, we discuss applications to animal movement studies and conservation initiatives.     

Spatial patterns of Mediterranean land abandonment and related land cover transitions

In Mediterranean France, land abandonment is a widespread change. To understand and predict the land abandonment process and its consequences, land cover change models are used. An essential step in the development of a land cover change model is the identification and quantification of the factors controlling land cover change. In this paper we present a change detection study using aerial photographs in combination with an extensive dataset of field data and geographical data, to identify and quantify these factors for a study area in Mediterranean France, 60 km west of the city of Montpellier. We distinguished 11 land cover change classes and 7 associated “time since abandonment” classes at a detailed scale. Several environmental and non-environmental factors were found to be important variables for the land abandonment process. Differences in soil class explain a large part of the land abandonment pattern and the associated transition paths and transition rates. Most abandoned lands are located on regosols and lithosols, which are marginal soils with respect to water holding capacity. Within soil classes, we could recognise different transition paths and transition rates. However, within the 55 years covered by this dataset detailed transitions from pioneer vegetation to vegetation higher in the succession, as described by other authors, were only found for a limited number of vegetation/soil combinations. We relate these slow transitions for some areas to ongoing grazing and for some other areas to irreversible degradation.     

Response of snails and slugs to fragmentation of lowland forests in NW Germany

Habitat fragmentation is a major cause for species loss, but its effect on invertebrates with low active dispersal power, like terrestrial gastropods, has rarely been studied. Such species can not cross a hostile habitat matrix, for which the predictions of island theory, such as positive relations between species richness and patch size, should apply. In order to test this prediction, we studied gastropod species diversity by assessing gastropod assemblage characteristics from 35 sites in 19 fragments of deciduous old-growth forests in the Lower Rhine Embayment, Germany. Assemblages differed between larger (≥700 ha) and smaller forests (<400 ha), those of large forests held a higher percentage of forest species. Although α-diversity was similar between the two forest size classes, small forests often comprised matrix species, resulting in a higher β-diversity. Edge effects on the species richness of matrix species were noticeable up to 250 m into the forest. Hierarchical partitioning revealed that distance to disturbances (external edge, internal edges like roads) explained most assemblage variables, whereas forest size and woodland cover within a 1 km radius from the sites explained only a few assemblage variables. Densities of two forest-associated species, Discus rotundatus and Arion fuscus, decreased with forest size. Yet, forest size was positively correlated with richness of typical forest species and densities of Limax cinereoniger. The latter species seems to need forests of >1,000 ha, i.e., well above the size of most fragments. In conclusion, the prediction is valid only for forest species. The response to fragmentation is species specific and seems to depend on habitat specialization and macroclimatic conditions. Jean-Pierre Maelfait: Deceased.     

Independent effects of connectivity predict homing success by northern flying squirrel in a forest mosaic

Landscape composition and configuration, often termed as habitat loss and fragmentation, are predicted to reduce species population viability, partly due to the restriction of movement in the landscape. Unfortunately, measuring the effects of habitat loss and fragmentation on functional connectivity is challenging because these variables are confounded, and often the motivation for movement by target species is unknown. Our objective was to determine the independent effects of landscape connectivity from the perspective of a mature forest specialist—the northern flying squirrel (Glaucomys sabrinus). To standardize movement motivation, we translocated 119 squirrels, at varying distances (0.18–3.8 km) from their home range across landscapes representing gradients in both habitat loss and fragmentation. We measured the physical connectedness of mature forest using an index of connectivity (landscape coincidence probability). Patches were considered connected if they were within the mean gliding distance of a flying squirrel. Homing success increased in landscapes with a higher connectivity index. However, homing time was not strongly predicted by habitat amount, connectivity index, or mean nearest neighbour and was best explained as a simple function of sex and distance translocated. Our study shows support for the independent effects of landscape configuration on animal movement at a spatial scale that encompasses several home ranges. We conclude that connectivity of mature forest should be considered for the conservation of some mature forest specialists, even in forest mosaics where the distinction between habitat and movement corridors are less distinct.     

Population structure and movements of freshwater turtles across a road-density gradient

Understanding interactions between roadways and population structure and movements of wildlife is key to mitigating “road effects” associated with increasing urbanization of the landscape. Aquatic turtles are a useful focal group because (1) population persistence is sensitive to mortality of individuals upon roads; (2) turtles frequently move among wetlands and encounter roads, and (3) turtles are an important component of vertebrate biomass in aquatic ecosystems. From 2005 to 2007, we examined the effects of urbanization on local- and landscape-scale populations of turtles. To do so, we sampled and marked turtles in 15 ponds arranged along a steep, urban–rural gradient in central New York State. We captured 494 turtles, representing 327 individuals, the majority of which were common snapping turtles Chelydra serpentina (n = 191) and eastern painted turtles Chrysemys picta picta (n = 122). At the local population (pond) scale, a higher proportion of female snapping turtles in ponds was associated with lower road densities within 500 m of ponds. The mean size of both species of turtle increased in ponds with a lower density of roads within 100 m. At the landscape-level, we observed fewer turtles dispersing through urbanized habitat than forested, and fewer movements through areas with a higher density of roads. Our study suggests that roads alter both local- and landscape-level turtle populations through a loss of female turtles, and by reducing movement between ponds. By extension, the study targets key landscape features upon which to focus mitigation efforts.     

Movement behaviour of a forest songbird in an urbanized landscape: the relative importance of patch-level effects and body condition during migratory stopover

With expansion of urban areas worldwide, migrating songbirds increasingly encounter fragmented landscapes where habitat patches are embedded in an urban matrix, yet how migrating birds respond to urbanization is poorly understood. Our research evaluated the relative importance of patch-level effects and body condition to movement behaviour of songbirds during migratory stopover within an urban landscape. We experimentally relocated 91 migrant Swainson’s thrushes (Catharus ustulatus) fitted with 0.66 g radio-transmitters to seven forest patches that differed in area (0.7–38.4 ha) and degree of urbanization within central Ohio, USA, May 2004–2007. Fine-scale movement rate of thrushes (n = 55) did not differ among urban forest sites, but birds in low energetic condition moved at higher rates, indicating an energetically mediated influence on movement behaviour. In larger sites, Swainson’s thrushes (n = 59) had greater coarse-level movement during the first 3 days and utilized areas farther from forest edge, indicating stronger influence by patch-level factors. Thrushes exhibited strong site tenacity, with only five individuals (7%) leaving release patches prior to migratory departure. Movement outside the release patch only occurred at the smallest forest patches (0.7 and 4.5 ha), suggesting that these sites were too small to meet needs of some individuals. Swainson’s thrushes exhibited edge avoidance and apparent area sensitivity within urban forest patches during stopover, implying that conservation of larger patches within urban and other fragmented landscapes may benefit this species and other migrant forest birds.     

Siberian flying squirrel responses to high- and low-contrast forest edges

We examined responses of Siberian flying squirrels ( Pteromys volans ) to edges between nesting habitat (mature spruce forests), movement habitat (other forests, pine bogs), and open areas within their home ranges in southern Finland in 1996-2000. Radio-tracked squirrels (n=146) were generally associated to edges when they were ac tive at night. Compared to distances expected from the habitat pattern of their home range, squirrels occurred closer to high-contrast edges (of open areas) and low-contrast edges (nesting or movement forest types). Asso ciation with edges of open areas was more pronounced when squirrels were in movement habitat than in nesting habitat, possibly because of stronger channeling of movements in the former habitat. When in nesting habitat, squirrels responded more strongly to field edges than to recent clearcut edges, probably as a result of the pres ence of more deciduous trees on field edges, unlike clearcut edges. Responses to open areas were independent of spatial scale. However, responses to movement habitat from nesting habitat, and vice versa, were more pronounced over hundreds than tens of meters. Nesting cavities and dreys were generally located at random with respect to edges. We conclude that squirrel responses to edges of landscape attributes are diverse and depend both on spatial scale and edge contrast.This revised version was published online in May 2005 with corrections to the Cover Date.     

Human pathways are barriers to beavers damaging trees and saplings in urban forests

Urban North American beaver (Castor canadensis) damage of trees and saplings was compared between shore forests and forests uphill of macadam, wood chip, and raised wood board human pathways used daily in Radnor Lake State Natural Area, Nashville, TN. Also, comparisons of beaver damage were made between shore forests and forests uphill of bare earth deer paths used less than once a month by humans and the forests were on 5% and 30% slopes. Means, standard deviations, and t-tests (P ≤ 0.05) were calculated for percent beaver damage, which included undamaged stems, beaver-cut stems, and beaver-cut stumps. Significant differences in beaver damage of trees and saplings were found between forests uphill of the human pathways used daily and the respective shore forests. Beaver damage of trees and saplings was not significantly different between the shore forests and forests uphill of the deer paths used less than once a month by humans for the 5% slope forest; however, the differences were significant for the 30% slope forest. Beaver damage of trees and saplings was significantly greater in the uphill of the deer paths forests than the uphill of the human pathways forests for comparable slope forests. Human scent on the pathways used daily made of macadam, wood chips, and raised wood boards was interpreted to be the barrier sensed by beavers to not cross over or under the human pathways to damage trees and saplings. This research suggests utilizing human pathways as an odor fence to spatially limit beaver damage, which provides a whole forest management alternative to individual tree protection for management of beaver damage in the urban forest.     

Predicting altered connectivity of patchy forests under group selection silviculture

Group selection silviculture creates canopy openings that can alter connectivity in patchy forests, thereby affecting wildlife movement and fire behavior. We examined effects of group selection silviculture on percolation (presence of continuously forested routes across a landscape) in Sierra Nevada East-side pine forest in northern California, USA. Four ~ 250 ha project areas were analyzed at three map resolutions in three ways: analyzing forest cover maps for percolation before and after group-selection treatment, placing simulated group openings in forest cover maps until fragmentation occurred, and comparing project areas to neutral maps that varied in forest cover and self-adjacency. Two project areas were fragmented (i.e., did not percolate) prior to treatment, one resisted fragmentation, and the other became fragmented by treatment when analyzed at 30 m cell resolution. Median simulated openings required to create fragmentation agreed well with the actual number. There was a well-defined transition between percolating and non-percolating neutral maps; increased aggregation of forest lowered the critical value at which forests percolated. A logistic model based on these maps predicted percolation behavior of the project areas effectively, but alternative generating algorithms gave slightly different predictions. A graph of this model provides a straightforward way to visualize how close a landscape is to fragmentation based on its forest cover and aggregation. In East-side Sierran landscape, fragmentation from group-selection openings may make the landscape less hospitable to the American marten but more resistant to crown fire.     

How key habitat features influence large terrestrial carnivore movements: waterholes and African lions in a semi-arid savanna of north-western Zimbabwe

Within a landscape where prey has an aggregated distribution, predators can take advantage of the spatial autocorrelation of prey density and intensify their search effort in areas of high prey density by using area-restricted search behaviour. In African arid and semi-arid savannas, large herbivores tend to aggregate around scarce water sources. We tested the hypothesis that water sources are a key determinant of habitat selection and movement patterns of large free-ranging predators in such savannas, using the example of the African lion. We used data from 19 GPS radio-collared lions in Hwange National Park, Zimbabwe. Maps of lions’ trajectories showed that waterholes are key loci on the lions’ route-maps. Compositional analyses revealed that lions significantly selected for areas located within 2 km of a waterhole. In addition, analysis of lions’ night paths showed that when lions are close to a waterhole (<2 km), they move at lower speed, cover shorter distances per night (both path length and net displacement) and follow a more tortuous path (higher turning angle, lower straightness index and higher fractal dimension) than when they are further from a waterhole. Hence, our results strongly suggest that lions adopt area-restricted searching in the vicinity of waterholes, and reduce their search effort to minimize the time spent far from a waterhole. They provide an illustration of how key habitat features that determine the dispersion of prey (e.g. waterholes in this study) have an influence on the spatial ecology and movement patterns of terrestrial predators.     

The utility of ancient forest indicator species in urban environments: A case study from Poznań, Poland

We aimed to assess the impact of land-use structure on AFIS occurrence and evaluation of these bioindicators in urban conditions. We compiled data about forest continuity based on archival maps and forest management plans; floristic records of 79 AFIS occurrences were collected within a 1 × 1 km grid and land-use form structure in Poznań (W Poland). We tested fidelity of AFIS using χ² tests and effects of land-use structure using random forest models. We also checked spatial autocorrelation and its impact on AFIS distribution patterns within old and recent forests, using spatially explicit generalized linear models. We found a strong relationship between AFIS number per grid square and fraction of forests and waters in land-use structure. Relationships between AFIS distribution and land-use shows that AFIS are a good proxy describing human impact or an urbanity gradient. AFIS number per grid square shows small, but significant positive spatial autocorrelation, which suggests possibilities of migration into new forests. AFIS, treated as a bioindicative group of species, may be a useful indicator of landscape and land-use structure transformation in urban environments. Occurrence of many AFIS that are endangered at the city scale indicates the need for conservation of old forests in urban areas, to preserve biodiversity in urban environments. Thus, AFIS may act as indicators of old forests with high conservation value in cities, which need special protection.     

Logging pattern and landscape changes over the last century at the boreal and deciduous forest transition in Eastern Canada

Forestry practices associated with the industrial era (since ~1900) have altered the natural disturbance regimes and greatly impacted the world’s forests. We quantified twentieth century logging patterns and regional scale consequences in three sub-boreal forest landscapes of Eastern Canada (117,000, 49,400 and 92,300 ha), comparing forestry maps depicting age and forest cover types for early industrial (1930) and present-day (2000) conditions. Results were similar for the three landscapes, indicating large-scale forest change during the twentieth century. In 1930, previous logging activities had been concentrated in the lowlands and along the main hydrographical network, as compared to a more even distribution over the landscapes in 2000, reflecting a decreasing influence of the environmental constraints on forest harvesting. In 1930, old-aged forests (>100 years) accounted for more than 75% of the unlogged areas of the three landscapes, as compared to less than 15% for the present-day conditions. Logging practices have thus inverted the stand age distribution of the landscapes that are currently dominated by young and regenerating stands. The 1930 forest cover types showed a clear relationship with elevation, with conifers located in the lowlands and mixed and deciduous stands restricted to the upper slopes. Between 1930 and 2000, 58–64% of the conifer areas transformed to mixed and deciduous forests, such that no clear altitudinal relationships remained in 2000. We conclude that twentieth century logging practices have strongly altered the preindustrial vegetation patterns in our study area, to the point that ecosystem-based management strategies should be developed to restore conifer dominance, altitudinal gradients, as well as the irregular structure inspired from old forest stands.     

Determining an appropriate minimum mapping unit in vegetation mapping for ecosystem restoration: a case study from the Everglades,USA

This paper documents the analyses that were conducted with regards to investigating an appropriate Minimum Mapping Unit (MMU) to be used to capture the potential changes in vegetation patterns for a 10,924 square km restoration project being conducted in south Florida, USA. Spatial landscape and class metrics that were shown to change predictably with increasing grain size were adopted from previous studies and applied to a multi-scale analysis. Specifically, this study examines the effects of changing grain size on landscape metrics, utilizing empirical data from a real landscape encompassing 234,913 ha of south Florida’s Everglades. The objective was to identify critical thresholds within landscape metrics, which can be used to provide insight in determining an appropriate MMU for vegetation mapping. Results from this study demonstrate that vegetation heterogeneity will exhibit dissimilar patterns when investigating the loss of information within landscape and class metrics, as grain size is increased. These results also support previous findings that suggest that landscape metric “scalograms” (the response curves of landscape metrics to changing grain size), are more likely to be successful for linking landscape pattern to ecological processes as both pattern and process in ecological systems often operate on multiple scales. This study also incorporates an economic cost for various grain dependant vegetation mapping scales. A final selection of the 50 × 50 m grain size for mapping vegetation was based on this study’s investigation of the “scalograms”, the costs, and a composite best professional judgment of seasoned scientists having extensive experience within these ecosystems.     

Intensive recreational activities in suburban forests: A method to quantify the reduction in timber value

Central European forests are generally multifunctional; the same forest areas are used for timber production and recreation. In frequently visited areas damage to trees can be observed. We developed a method to estimate the actual reduction in timber value due to recreation-induced damage. The method was tested in two suburban (oak–hornbeam and beech) forests sustainably maintained by a selective management system and with free access in northwestern Switzerland. Considering the total forest areas, 9.4% of oak trees and 23.0% of beech trees were damaged by recreational activities. The resulting reduction in timber value averaged 19 and 53 € ha^?1 a^?1 in the two forests. The annual reduction in timber value due to recreation-induced damage can account for up to 16% of the total proceeds. The monetary benefits of forest recreation in these areas, however, by far exceed the damage to trees.